JACKSONVILLE – The Jaguars cut their roster to 53 Saturday.

That had to be done to comply with NFL rules, but what David Caldwell on Saturday called the most difficult "final roster" moves in his three seasons as the Jaguars' general manager may not yet actually be final.

The Jaguars on Saturday not only released safety Craig Loston, offensive lineman Austin Pasztor and linebacker Jeremiah George, they also placed defensive tackle Ziggy Hood on injured reserve. The moves trimmed the roster to 53 players.

All NFL teams had until 4 p.m. Eastern Saturday to reach that number.

"This was a tough one," Caldwell said of the 22 roster moves the team made Friday and Saturday to reach the 53-player limit. "In the last couple of years, we were able to make them all in one fell swoop. This year, we had to wait on a few and finagle some things with injured reserve and PUP."

Caldwell said the Jaguars already had scouted players they believed could be released around the league Saturday.

"We have a handful of guys we're working through," Caldwell said. "We just have to make sure if we make a move it is to make us get better and it's not for the sake of making a move to make a move. It depends on what comes available. We have to wait and see what comes available."

The Jaguars also on Saturday moved defensive tackle Sen'Derrick Marks from the physically unable to perform list to the active roster, meaning Marks is on the 53-player roster. Marks spent the offseason rehabilitating after January reconstructive knee surgery.

Caldwell said Marks is not expected to play in the regular-season opener against Carolina Sunday, adding that he believes Marks could play before Week 6. Players kept on the PUP list to start the regular season can't play or begin practicing until after Week 6.

"Once he starts to practice I think we'll have a better idea on that," Caldwell said, adding that Marks could begin doing individual drills during practice leading to the team's Week 2 game against Miami.

Hood was placed on injured reserve after sustaining a plantar fasciitis injury in the preseason finale at Washington Thursday. With Hood out and Marks unavailable, Caldwell said Tyson Alualu will play Marks' three-technique tackle position along with end Jared Odrick, rookie Michael Bennett and veteran reserve Abry Jones.

Caldwell said the decision to release Loston – a second-year veteran who played 13 games last season – and keep third-year veteran Josh Evans had to do with Evans' experience and Loston's health. Evans has played free and strong safety in his first two seasons with the Jaguars and Loston sustained an ankle injury in the preseason finale at Washington Thursday.

The Jaguars kept four safeties on the 53: strong safety Johnathan Cyprien, who Caldwell said is expected to play in the opener after missing the final three preseason games with a finger injury; free safety Sergio Brown; Evans; and rookie safety James Sample, who played only in the preseason finale after missing the first three preseason games with a broken arm.

Caldwell said Cyprien likely will play Week 1 with a cast on the base of his finger.

"Our doctors passed him," Caldwell said of Loston, "but the big thing was Josh knowing both positions and Sample being a rookie. Not knowing if Craig would be 100 percent or not – we thought he would be OK for practice, but there was just a concern there."

Caldwell also addressed the status of defensive end Andre Branch, saying the four-year veteran could be ready by perhaps Week 4. He sustained a sprained medial collateral ligament in Preseason Week 3.

"He'll be a situation we reevaluate on a week-to-week basis and once we get him out there on the field we should know a little better timeframe," Caldwell said.

The Jaguars on Friday released 18 players: quarterback Stephen Morris, cornerback Jeremy Harris, running back Storm Johnson, wide receiver Neal Sterling, tight end Ben Koyack, defensive tackle Richard Ash, defensive end Camaron Beard, defensive end Cap Capi, wide receiver Kasey Closs, offensive lineman Will Corbin, tight end Connor Hamlett, defensive end Ikponmwosa Igbinosun, wide receiver Erik Lora, offensive guard Chris Reed, cornerback Rashaad Reynolds, linebacker Todd Thomas, cornerback Peyton Thompson and wide receiver Tony Washington.

Teams can start signing players to their 10-player practice squads on Sunday.

"We let some of these know that would be a possibility," Caldwell said.

Caldwell also on Saturday:

*Said second-year wide receiver Marqise Lee is considered about "50-50" to play against the Panthers. He has been out since early in training camp with a hamstring injury. "If we hold him out opening week it's probably more of a precaution so he can hit the ground running Week 2," Caldwell said. …

*Updated the status of tight end Julius Thomas, saying the free-agent signee had a screw placed in his hand in surgery Thursday. Caldwell said Thomas will be reevaluated in two weeks. "I think we'll have a better timeframe to see how he's responding at that point in time," Caldwell said. …

*Discussed veteran John Lotulelei, who made the roster as a reserve linebacker and special teams player because of what Caldwell called "his ability to disrupt and fly around." Caldwell added, "He can go out and play at a very high pace with a lot of urgency and speed and create things. He may have come in to this camp as a little bit of a long shot but he has done enough to win a job." …

*Discussed rookie cornerback Nick Marshall, who made the roster after signing shortly after the 2015 NFL Draft as a collegiate free agent. He played quarterback for Auburn the past two seasons, moving to cornerback at the Senior Bowl in January. "We want to try to get guys who can really make a difference on special teams for us," Caldwell said, adding that Marshall has backup returner ability and can cover kickoffs. "In terms of him playing corner, when you get down to your fifth corner on game day, it's probably not good for anybody in the league. We said, 'This is a guy who has length, speed and athleticism.' We feel like there's a lot of upside for him, but it really came down to special teams."